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Toshio Shibuya, Ryosuke Endo, Yoshiaki Kitaya, and Saki Hayashi

with the light interception area per plant and the net photosynthetic rate per unit leaf area, respectively. The high-R:FR light has been shown to alter these growth parameters. There have been many reports that leaf enlargement (correlated with LAR

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Daniel J. Barta and Theodore W. Tibbitts

Tissue concentrations of Ca, Mg, and K were determined across immature leaves of lettuce (Lactuca sativa L. `Buttercrunch') at different stages of enlargement using electron microprobe x-ray analysis. The analysis was with a wavelength dispersive spectrometer to permit detection of low concentrations of Ca. Patterns of mineral accumulation in immature leaves that were exposed were compared to patterns of accumulation in leaves that were enclosed within a developing head. The leaves developing without enclosure were free to transpire and developed normally whereas leaves developing with enclosure were restricted in transpiration and developed an injury that was characteristic of Ca deficiency. In the exposed leaves, Ca concentrations increased from an average of 1.0 to 2.1 mg·g-1 dry weight (DW) as the leaves enlarged from 5 to 30 mm in length. In the enclosed leaves, Ca concentrations decreased from 1.0 to 0.7 mg·g-1 DW as the leaves enlarged from 5 to 30 mm in length. At the tips of these enclosed leaves a larger decrease was found, from 0.9 to 0.3 mg·g-1 DW during enlargement. Necrotic injury first became apparent in this tip area when the concentration was ≈0.4 mg·g-1 DW. Magnesium concentrations across the exposed leaves were similar to concentrations across the enclosed leaves, and did not change with enlargement. Magnesium concentrations averaged 3.5. mg·g-1 DW in both enclosed and exposed leaves during enlargement from 5 to 30 mm. In both exposed and enclosed leaves, K concentrations increased during enlargement from 40 to ≈60 mg·g-1 DW. Potassium concentrations were highest toward the leaf apex and upper margin where injury symptoms occurred, and this may have enhanced injury development. This research documents the critical low levels of Ca (0.2 to 0.4 mg·g-1 DW) that can occur in enclosed leaves of plants and which apparently leads to the marginal apex necrosis of developing leaves seen frequently on lettuce and other crops.

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R. Bruce Carle and J. Brent Loy

The morphology, growth rate and anatomy of the fused vein trait were characterized in Cucurbita pepo using the inbreds NH2405 (fused vein), NH7210 (moderately fused vein), and NH614 (normal). Morphological analysis showed that the trait is characterized by a partial fusion of the five primary leaf veins. Fusion begins at the distal point of the petiole and extends along the central vein. Branching of the veins is delayed and there is a reduction of the interveinal leaf blade. Consequently, the upper leaf surface appears puckered or wrinkled. Depending on genetic background, the onset of fused vein leaf production starts at the fourth to tenth leaf stage and continues throughout vegetative growth. The extent of fusion increases with leaf number but stabilizes by the twentieth leaf stage maximum extent of vein fusion also varies with genetic background (5-20 cm). Though fused vein and normal inbreds differed in the rate and pattern of leaf growth, examination of F2 and BC populations revealed no significant effect of the fused vein trait on leaf number, leaf size, and rate of leaf initiation. Anatomical examination revealed different vascular patterns in the transition zone between petiole and leaf blade for normal and fused vein leaves. In normal leaves, the vascular bundles of the petiole enlarge and coalesce to form a vascular crescent. The crescent reorganizes and diverges as large vascular columns and pairs of smaller flanking vascular bundles into each vein. In contrast, two cycles of enlargement, coalescence, and dispersal occur in fused vein leaves.

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W.L. Berry, R.M. Wheeler, C.L. Mackowiak, G.W. Stutte, and J.C. Sager

Critical levels of nutrients in leaf tissue are influenced by plant metabolism, environment, and nutrient availability. In this study, we measured the elemental concentrations in fully expanded, upper canopy potato (Solunum tuberosum cv. Norland) leaves throughout growth and development in a controlled environment. Plants were grown hydroponically (NFT) in NASA's Biomass Production Chamber using a complete nutrient solution with the electrical conductivity maintained continuously at 0.12 S m-1. Photoperiod and air and root zone temperatures were changed midseason to promote tuberization, while CO2 levels were maintained at 1000 μmol mol-1 throughout growth. During vegetative growth, leaf nutrient concentrations remained relatively constant, except for a decline in Ca. During tuber enlargement and plant maturation, overall nutrient uptake decreased. Concentrations of the less mobile nutrients such as Ca, Mg, and B increased in the leaf tissue during mature growth, but somewhat surprisingly, highly mobile K also increased. Leaf concentrations of P, Zn, and Cu decreased during maturation.

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D.S. Achor and L.G. Albrigo

Permanent chlorosis of leaves on plants fertilized with urea containing high levels of the contaminant biuret has been observed in several crops including citrus. Little has been reported as to the cellular changes that result from such chlorosis. Branches from `Ruby Red' grapefruit (Citrus paradisi Macfadyn) and `Hamlin' orange [C. sinensis (L.) Osbeck] were sprayed with urea solutions containing 1.05% biuret. As visible symptoms developed, leaf tissue samples were prepared for transmission electron microscopy. For comparison purposes, leaves from similar trees showing chlorosis from age-related senescence and Zn deficiency were also sampled. The progressive development of chlorosis in biuret-affected leaves was characterized by: the loss of starch, thylakoidal and granal membranes in chloroplasts along with the enlargement and increase in number of plastoglobuli or lipid bodies. The lipid bodies were liberated alone or in association with membrane vesicles to the cytoplasm and vacuoles. The number and volume of the individual chloroplasts became smaller. Concurrent loss of cytoplasmic content and the enlargement of the vacuolar space were also observed in the biuret affected leaf tissue. Similar findings were observed in the cells of senescent leaves. In cells of leaves showing nutritional deficiency, losses in cytoplasmic content and vacuolar enlargement were observed but there was neither complete loss of thylakoidal or granal membranes nor the release of lipids from the plastids. It was concluded that 1) the cytological characteristics of the biuret-affected samples were more similar to age-related senescent samples than to chlorosis from Zn deficiency and 2) that complete loss of the lipid bodies from the chromoplasts to the cytoplasm and vacuole in the biuret-affected samples and in age-related senescence in citrus leaves was responsible for the permanent nature of the chlorosis.

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Rongcai Yuan, Fernando Alferez, Igor Kostenyuk, Shila Singh, James P. Syvertsen, and Jacqueline K. Burns

The effects of 2 consecutive years of annual defoliation during the harvest season on fruit size, yield, juice quality, leaf size and number were examined in trees of the midseason cultivar `Hamlin' and the late-season cultivar `Valencia' orange [Citrus sinensis (L.) Osb.]. In `Hamlin', removal of up to 50% of the leaves in late November had no effect on fruit yield, fruit number, fruit size, soluble solids yield, juice °Brix, and °Brix to acid ratio of juice the following year. In `Valencia', removal of 50% of the leaves in late March decreased fruit yield and soluble solids yield but did not affect Brix or the Brix to acid ratio of the juice. Leaf size of new flush was reduced by removal of 50% of the leaves in both cultivars but there was little effect on total canopy size. There were no measured effects of removing 25% of leaves from tree canopies. Thus, canopy growth, fruit yield, fruit quality, and leaf size were not negatively impacted when annual defoliations did not exceed 25% of the total canopy leaf area in `Valencia' and `Hamlin' orange trees for two consecutive years. Overall, fruit weight increased linearly with increasing ratio of leaf area to fruit number, suggesting that fruit enlargement can be limited by leaf area.

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Rongcai Yuan, Francisco Garcia-Sanchez, Fernando Alferez, Igor Kostenyuk, Shila Singh, Guangyan Zhong, James Syvertsen, and Jacqueline Burns

The effect of annual defoliation over two consecutive years on fruit yield, juice quality, leaf size, and number was examined in 11-year-old `Hamlin' and 13-year-old `Valencia' orange [Citrus sinensis (L.) Osb.] trees. Removal of up to 50% of the leaves in late November had no effect on fruit number, fruit weight, fruit yield, soluble solids yield, juice °Brix, and °Brix: acid ratio of juice in `Hamlin' oranges. In `Valencia' oranges, removal of up to 50% of the leaves in late March also did not affect °Brix or the °Brix: acid ratio of the juice, but decreased fruit yield and soluble solids yield. Leaf size was reduced by removal of 50% of the leaves in both cultivars. Removal of up to 50% leaves in late November had no significant influence on net CO2 assimilation (aCO2) of the subsequent spring flush leaves in early May in `Hamlin' oranges, whereas aCO2 of `Valencia' spring flush leaves in early May increased linearly with increasing levels of defoliation in late March. The results indicate that fruit yield, fruit quality, leaf size, and number were not negatively impacted when annual defoliations did not exceed 25% of the total canopy leaf area for `Valencia' and `Hamlin' oranges for two consecutive years. Overall, in whole `Hamlin' or `Valencia' orange trees, fruit weight increased linearly with increasing ratio of leaf area to fruit, suggesting that fruit enlargement depends on available photosynthate and can be limited by leaf area.

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Cary A. Mitchel

Brief, periodic seismic (shaking) or thigmic (contact rubbing) stress treatments applied to plants growing in a wind-protected environment typically reduce but strengthen vegetative growth and often inhibit reproductive development. Cell division and cell enlargement both are affected. Mechanically dwarfed plants accumulate less leaf area than do undisturbed controls and undergo temporary stomatal aperture reduction following an episode of stress, leading to reduced photosynthetic productivity. Vibration or mild shaking may lead to a slight stimulation of plant growth. Most classes of phytohormones have been implicated to mediate different growth responses to mechanical stress. Physical perturbation turns on the transcription of several genes coding for calmodulin-like proteins. Calcium chelators and calmodulin inhibitors partially negate effects of thigmic stress. Growth rate responses of naive seedlings are immediate and dramatic, suggesting turgor collapse, whereas recoveries are slow and sometimes partial, suggesting reduced wall extensibility in the cell enlargement zone. Mechanical stress may be used for height control of intensively cultivated bench crops or to physically toughen bedding plants prior to outdoor transplant. Physiological hardening remains a question. Mechanical height control avoids use of chemicals but increases risk of wounding and pathogen infection.

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Theodore C. Hsiao

Of all the plant processes examined, leaf growth and canopy development is the most sensitive to water stress. The consequent reduction in cumulative radiation interception by the plant leads to a smaller biomass as well as reduced transpiration, usually without altering radiation-use efficiency or water-use efficiency of the canopy. Sensitivity of leaf growth to the growth medium or aerial environment of the plant will be illustrated. A way to quantify the consequent and often marked impact on productivity will be discussed. In contrast with the high sensitivity of leaf growth to water stress, root growth is more resistant. This allows at least the partial maintenance of root growth as the stress intensifies. The result is a more thorough extraction of soil water while transpiration is restricted by the smaller leaf area. The possible mechanisms for the differential sensitivity of leaf and root growth to water stress will be evaluated. Emphasis will be placed on processes underlying cell enlargement. Recent data, obtained with the pressure microprobe that measures turgor pressure in individual cells, will be presented to illustrate the contrasting responses in growth, cell wall extending ability, and solute transport to the growing cells when the plant adjusts and accommodates to changes in water status.

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Karen I. Theron and Gerard Jacobs

Flowering-size Nerine bowdenii bulbs were sampled from a commercial planting at 2-week intervals from 13 Aug. 1991 to 14 June 1992. They were dissected, and the following variables were recorded: 1) number and dry weight of fully sheathing leaf bases or leaves of each growth unit, 2) length and dry weight of foliage leaves, 3) fresh weight of outermost inflorescence, and 4) dry weight of daughter bulbs. Bulb organs that served as sinks and sources changed as the bulb progressed in its growth and developmental cycle. Before the new foliage provided photosynthates, growth depended on reserves deposited and stored in leaf bases during the preceding seasons. Reserves were used for the development of new leaves (foliage and bases), roots, and daughter bulb enlargement. Once the foliage became the photosynthate source, reserves were stored in old and new leaf bases. The inflorescence became the major sink when elongation of the scape initiated. Thereafter, daughter bulbs became the dominant sinks, receiving photosynthates from the senescence foliage and the reserves stored in leaf bases. The decrease in dry weight of the leaf bases was prominent in bulbs that remained in situ.